Research Article |
Corresponding author: Ângelo Parise Pinto ( odonata_angelo@hotmail.com ) Academic editor: Gabriel L. F. Mejdalani
© 2018 Ângelo Parise Pinto, Tom Kompier.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pinto ÂP, Kompier T (2018) In honor of conservation of the Brazilian Atlantic Forest: description of two new damselflies of the genus Forcepsioneura discovered in private protected areas (Odonata: Coenagrionidae). Zoologia 35: 1-19. https://doi.org/10.3897/zoologia.35.e21351
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Two new Brazilian Protoneurinae damselflies, Forcepsioneura regua sp. nov. (holotype male deposited in DZRJ: Brazil, Rio de Janeiro State, Cachoeiras de Macacu municipality, RPPN Reserva Ecológica de Guapiaçu) and Forcepsioneura serrabonita sp. nov. (holotype male deposited in DZRJ: Brazil, Bahia State, Camacan municipality, RPPNs Serra Bonita) , are described, illustrated and diagnosed based on males and females. The bluish and smaller F. regua sp. nov. has been confused with at least three previously described species, being very similar to the type species of Forcepsioneura, F. garrisoni Lencioni, 1999, but lacking a defined tubercle-like process on the posterolateral margin of the median lobe of the prothorax in both sexes, which allows it to be distinguished from all other known species. The shape of the cercus of the male of F. serrabonita sp. nov. is similar to that of F. grossiorum Machado, 2001 and F. lucia Machado, 2000, two species with very short ventrobasal process. However, it differs from them mainly by the mediobasal process of the cercus, which is rounded in dorsal view and almost not visible in lateral view. The taxonomic status of Forcepsioneura is discussed and a comparison with the other species of the genus is provided. Based on size, habitat and coloration, Forcepsioneura can be informally divided into two groups: (1) large, orange-black and montane species, including F. grossiorum, F. itatiaiae (Santos, 1970), F. lucia and F. serrabonita sp. nov.; (2) small, bluish and lowland species, including F. garrisoni, F. haerteli Machado, 2001, F. regua sp. nov. and F. sancta (Hagen in Selys, 1860). Our findings highlight the urgency in directing collecting efforts to unexplored areas, as well as the importance of private preserves that harbor the type localities as guardians of the threatened and diverse Atlantic Forest diversity. Together these two localities surveyed account for more than 210 species of odonates, representing almost 24% of the number of Brazilian species. Brazil has the greatest number of known species of odonates in the world. This study shows that further research is required in order to fully understand the diversity of Forcepsioneura.
Amazoneura , dragonfly, damselfly, Protoneurinae , Southeastern Brazil, taxonomy
“…forest cover must be important for adults of the majority of neotropical Odonata. Every effort should be made to preserve forests and forest wetlands in this most biodiverse region of the Earth” (
The quotation above illustrates the relevance of forests for the conservation of the insects of the order Odonata, popularly known as dragonflies and damselflies. Life on Earth is facing the sixth mass extinction in the planet’s geological history (
Amongst the severely threatened areas of the biosphere, the Atlantic Forest (AF) is recognized as an area of immense species diversity and, for some groups of plants, such as Bromeliaceae, the richest one (
Protoneuridae s.l., as defined by
South American protoneurids are mostly small sized odonates living near or within forested areas. They are generally weak fliers, frequenting shaded areas near the shoreline of large freshwater bodies and numerous running water environments. Because of very thin general aspect and disruptive colors, protoneurids are difficult to be found in the field, being often overlooked by regular collectors.
Today’s Brazilian fauna counts with 70 species of Protoneurinae in 12 genera (
Forcepsioneura belongs to a poorly-supported monophyletic group of six genera, along with Amazoneura Machado, 2004, Lamproneura De Marmels, 2003, Roppaneura Santos, 1966, Phasmoneura Williamson, 1916, and Psaironeura Williamson, 1915 (
From the six species currently recognized in Forcepsioneura, only F. sancta was described a long time ago. Most species were discovered recently, thanks to field-works in poor explored areas, and undescribed species are often reported in the literature (e.g.,
In recognition of conservation efforts to protect AF remnants, we describe here two previously unknown damselflies of “Protoneuridae s.s.” (Coenagrionidae: Protoneurinae) recently collected in the states of Bahia and Rio de Janeiro, Brazil. These new damselflies were discovered in protected areas maintained thanks to private associations. The Brazilian AF is one of the most threatened biotic domains of the biosphere, and given its fragmentation and fragility, all conservation actions, including private sector initiatives, must be applauded for their contribution to both the scientific community and society in general.
Specimens examined are deposited in the following collections. Acronyms, wherever possible, based on the updated version of Arnett et al. (
General morphology terms follow
All measurements (in mm) and illustrations were made with the aid of a stereomicroscope equipped with a camera lucida. Specimens were photographed with a Leica MZ16 stereomicroscope equipped with a Leica DFC420 camera, and source images combined using Auto-montage software by The Synoptic Group©.
The following abbreviations were used in the text: a.s.l. = above sea level; Ax = antenodal crossveins; Fw = fore wing; GL = genital ligula; Hw = hind wing; MBP = mediobasal process; Px = postnodal crossveins; Pt = pterostigma; S1–10 = abdominal segments; L1–2 = segments of genital ligula; VBP = ventrobasal process.
Forcepsioneura
garrisoni
nec Lencioni, 1999: —
Forcepsioneura
haerteli
nec Machado, 2001: —
Forcepsioneura
sp.: —
Small, dorsally metallic greenish-blue, lateroventrally light blue and yellowish striped protoneurid; frons angulated; rear of head pale; CuP&AA indistinct; Ax space 1 as long as Ax space 3; GL with long inner fold and distal lateral lobe prolonged into a flagellum; cercus forcipate with mediobasal (MBP) and ventrobasal (VBP) processes, except for ill-defined prothoracic tubercle-like process, all characters agreeing with the definition of Forcepsioneura as recently diagnosed (Machado 2009,
The comparatively longer VBP, longer than 0.53 of total cercus length (dorsal branch) in lateral view, allows separation of males of F. regua sp. nov. from species of the orange-black group (VBP ≤ 0.40 of cercus length in F. grossiorum, F. lucia, and F. serrabonita sp. nov.); anteromesal margin of MBP acute in dorsal view (Fig.
Based on its general color and size, F. regua sp. nov. is most similar to F. garrisoni. Indeed, these two species are remarkably similar due to the blue stripes on thorax (light blue group). The short MP vein, which reaches distally at most 0.5 of the length of cell distal to the vein descending from subnodus, and the fin-shaped plate of MBP with an acute and strongly anteriorly directed apex in laterodorsal view are unique to these two species and separate them from all other known species in the genus. The short VBP, not reaching ventral margin of S10 ventrally, and posterolateral margin of median lobe of prothorax lacking a defined tubercle-like process distinguish F. regua sp. nov. from F. garrisoni (VBP long, ventrally reaching the ventral margin of S10, and posterolateral margin of median lobe of prothorax with a strongly defined conical tubercle-like projection). The ill-defined process on prothorax is unique for F. regua sp. nov. (Figs
Despite the fact that the blue coloration and the ill-defined process on the prothorax of F. regua sp. nov. appear to be unique to females of this species, females of other Forcepsioneura are poorly known. Consequently, females should be identified through comparison with descriptions, figures, and their association with males.
Head (Figs
Thorax (Figs
Wings (Fig.
Abdomen (Figs
Measurements (mm). Total length (incl. caudal appendages) 33.5; abdomen length (excluding caudal appendages) 28.8; head maximum width 2.9; Fw length 18.5; Hw length 17.4; Fw maximum width 3.3, Hw maximum width 3.3; Pt length 0.47 in Fw, 0.49 in Hw; length of metathoracic femur 1.6; metathoracic tibia 1.7; length of S9+10 in lateral view 1.1; length of cercus (dorsal branch) in lateral view 0.45; length of ventrobasal process in lateral view 0.29.
Similar to holotype but paler, differences described below.
Head (Fig.
Thorax (Figs
Abdomen (Figs
Measurements (mm). Total length (incl. caudal appendages) 35.2; abdomen length (excluding caudal appendages) 30.5; head maximum width 2.9; Fw length 21.2; Hw length 20.2; Fw maximum width 3.7; Hw maximum width 3.8; Pt length 0.53 in Fw; 0.54 in Hw; length of metathoracic femur 2.0; metathoracic tibia 2.2; length of S9+10 in lateral view 1.0; total length of cercus in lateral view 0.28.
The nine other males are very similar to holotype except for minor differences described below.
Head: Antefrons ivory yellow with a mesal ill-defined brown line; occipital bar with a yellow spot. Flagellum dark brown. Thorax: Prothorax paler with whitish-blue ventrolateral stripe faint; posterior lobe truncated or with small median concavity. Light blue longitudinal stripe on synthorax occupying at maximum 0.5 of mesepimeron width to almost entire metepisternum. Long and robust spurs on anteroventral surface of femora 3–4 in pro-, 4 in meso- and 4–5 in meta-, posteroventral surface with shorter and thinner spurs, 3–5 in pro-, 4–6 in meso- and 5–8 in metathoracic femora; anteroventral surface of tibiae armed with 9–11 spurs in pro- (4–5 of tibial comb), 5–6 in meso-, 5–6 in metathorax.
Abdomen: Paraproct orange-yellow to dark brown.
Measurements (mm, n = 5). Total length (incl. caudal appendages) 31.7–35.9; abdomen length (excluding caudal appendages) 27.0–30.8; head maximum width 2.8–3.0; Fw length 17.3–19.2; Hw length 16.5–18.0; Fw maximum width 3.0–3.5; Hw maximum width 2.9–3.3; Pt length 0.44–0.58 in Fw; 0.46–0.58 in Hw; length of metathoracic femur 1.7–1.9; metathoracic tibia 1.9–2.1; length of S9+10 in lateral view 1.1–1.3; length of cercus (dorsal branch) in lateral view 0.42–0.60; length of ventrobasal process in lateral view 0.30–0.36.
Unknown.
(10 males, 2 females). Holotype male and paratype female (in tandem), BRAZIL. Rio de Janeiro State: Cachoeiras de Macacu municipality, Reserva Ecológica de Guapiaçu (REGUA), forest fragment (22°28’08”S, 42°45’39”W, 42 m a.s.l.), 27.VII.2012, T.M.F. Kompier leg. (DZRJ 2071); 1 male paratype, same data but pond with macrophytes (22°27’10”S, 42°46’13”W, 34 m a.s.l.), 03.XII.2009, A.L. Carvalho & Disciplina Técnicas de Coleta PPGZoo/UFRJ leg. (DZRJ 315, DNA voucher ENT3609); 1 male paratype, same data but stream at small wetland (22°25’51”S, 42°45’39”W, 75 m a.s.l.), 02.III.2013, T.M.F. Kompier leg. (DZRJ 326); 2 male paratypes, same data but 22°28’04”S, 42°45’32”W, 70 m a.s.l., 13.I.2014, (DZRJ 2251, DNA voucher ENT2854; DZRJ 2252, DNA voucher ENT3608); 1 male paratype, same municipality but [District of] Japuíba [22°33’46”S, 42°41’30”W, 26 m a.s.l.], Rio Santana (sic, Rio Macacu?), 08.V.1983, J.M. Costa leg. (MNRJ ODO-0011, DNA voucher ENT2368); 2 male paratypes, Mangaratiba municipality, Ilha de Marambaia, Gruta da Santa (23°03’35”S, 43°57’56”W, 100 m a.s.l.), 02.XII.2000, R.W. Garrison leg. (RWG, DNA voucher ENT3016; DZUP 498845); 1 male and 1 female paratypes, same data but 04.XII.2000 (RWG); 1 male paratype, same data but temporary pool close to areal (MNRJ-ODO 0208, DNA voucher ENT3607). Specimens in DZRJ collected under ICMBIO/SISBIO license numbers 25034-1, 25034-2, and 25034-3.
Males were observed at seeps from peaty substrate at the forest edge and on very shallow, small streams with abundant leaf litter and limited flow. Even though observations were made on sunny days, they seem to avoid direct sunlight, perching suspended from the tips of leaves and grasses, or on the leaf litter. Streams were either partially or fully covered by trees and bushes. Females were observed in the same habitat. Both males and females were seen flying very slowly, and appeared to float in the air. Copulation lasted up to 10 minutes. Females were observed ovipositing unattended on a muddy substrate. Their habitat was shared with Heteragrion aurantiacum Selys, 1862, H. consors Hagen in Selys, 1862 and Acanthagrion gracile (Rambur, 1842), species that are tolerant to disturbed habitats in typical southeastern Brazilian Atlantic Forest.
Noun in apposition in reference to the acronym of the type locality, Reserva Ecológica de Guapiaçu (REGUA), a private reserve maintained for conservation, scientific research and environmental education.
This species has been known to specialists for at least eighteen years (e.g.,
Prothorax and genital ligula of males of Forcepsioneura: (13–16) F. serrabonita sp. nov., holotype (Brazil. BA: Camacan, DZRJ 0050); (17–20) F. regua sp. nov., (17) holotype (Brazil. RJ: Cachoeiras de Macacu, DZRJ 2071), remainder from paratype of same locality; (13, 17) prothorax in dorsal view; (14–16, 18–20) genital ligula in ventral (14, 18) and lateral (15–16, 19–20) views. Scale bars = 0.5 mm.
Caudal appendages of males of Forcepsioneura: (21–23) F. serrabonita sp. nov., holotype (Brazil. BA: Camacan, DZRJ 0050); (24–26) F. regua sp. nov., holotype (Brazil. RJ: Cachoeiras de Macacu, DZRJ 2071); (21, 24) lateral view; (22, 25) dorsal view; (23, 26) posterior view. Scale bars = 1 mm.
A medium, dorsally black and lateroventrally yellowish protoneurid species; frons angulated; rear of head pale; posterolateral margin of median lobe of prothorax with well-defined tubercle-like process; CuP&AA indistinct; Ax space 1 as long as Ax space 3; GL with long inner fold and distal lateral lobe prolonged into a flagellum; cercus forcipate with mediobasal (MBP) and ventrobasal process (VBP), all characters agreeing with the definition of Forcepsioneura as recently diagnosed (Machado 2009,
Based on general color and shape of male cercus, F. serrabonita sp. nov. resembles F. grossiorum and F. lucia. Males of this orange-black group of large species with robust cercus have a short VBP (as long as ≤ 0.40 of dorsal branch of cercus in lateral view), while it is longer (as long as ≥ 0.55 of dorsal branch) in F. garrisoni, F. haerteli, F. itatiaiae, F. regua sp. nov. and F. sancta. Forcepsioneura serrabonita sp. nov. is distinguished from the first two species by having posterior lobe of prothorax regularly rounded (Figs
In view of the large size of specimens of F. serrabonita sp. nov., they can be confused with F. itatiaiae, from which, in addition to the shorter VBP mentioned above, it can be separated by the shape of the male cercus with strongly cylindrical VBP (Figs
Additionally, the distinctive shape of the GL (Figs
Females of several Forcepsioneura species are poorly known, which hampers a useful diagnosis. Putative females should be identified by comparison with descriptions, figures and association with males.
Head (Figs
Thorax (Figs
Wings (Fig.
Abdomen (Figs
Measurements (mm). Total length (incl. caudal appendages) 38.7; abdomen length (excluding caudal appendages) 33.0; head maximum width 3.4; Fw length 21.8; Hw length 20.3; Fw maximum width 3.8, 3.7 in Hw; Pt length 0.54 in Fw, 0.58 in Hw; length of metathoracic femur 2.0; metathoracic tibia 2.1; length of S9+10 in lateral view 1.46; total length of cercus (dorsal branch) in lateral view 0.88; length of ventrobasal process in lateral view 0.36.
Similar to holotype. The only differences are described below.
Head (Figs
Thorax (Figs
Wings: MP reaches anal margin slightly distal of the vein descending from subnodus, covering 2 cells and ca 0.25 of a third cell in FW and 0.5 in HW. Px in Fw 13; Hw 11; RP2 originating at or slightly proximal to Px 6 in Fw, at Px 4 in Hw.
Abdomen (Figs
Measurements (mm). Total length (incl. caudal appendages) 37.3; abdomen length (excluding caudal appendages) 31.0; head maximum width 3.1; Fw length 21.2; Hw length 19.8; Fw maximum width 3.8; Hw maximum width 3.9; Pt length in Fw 0.54; Pt length in Hw 0.59; length of metathoracic femur 2.2; metathoracic tibia 2.4; length of S9+10 in lateral view 1.0; total length of cercus (dorsal branch) in lateral view 0.32.
The four other specimens (excluding the teneral one) are very similar to holotype, minor differences are described below.
Head: Labrum brownish to black. Postclypeus shining dark or with ill-defined mesal orange-brown spot. Frons and remainder of epicranium opaque black with a weak copper luster. Antennal flagellum brown to black. Posterior region of cranium (“postgena” plus “occiput”) ivory yellow.
Thorax: Anteroventral surface of femora armed with long and robust spurs, 5 in pro-, 3–4 in meso- and 5 in metathoracic femora, posteroventral surface with shorter and thinner spurs, 3 in pro-, 4–5 in meso-, 5–6 in metathoracic femora; anteroventral surface of tibiae armed with 9–10 spurs in pro- (4–5 of tibial comb), 5–6 in meso-, 5–6 in metathoracic tibiae.
Wings: MP reaches anal margin at or slightly distally to vein descending from subnodus, covering from 0.33 to 0.5 of a third cell in HW. Px in Fw 12–15; Hw 10–13; RP2 originating between Px 5–6 in Fw.
Measurements (mm). Total length (incl. caudal appendages) 37.5–40.5; abdomen length (excluding caudal appendages) 31.7–33.7; head maximum width 2.9–3.3; Fw length 20.3–22.3; Hw length 19.5–21.3; Fw maximum width 3.5–3.8; Hw maximum width 3.7–3.8; Pt length 0.56–0.60 in Fw; 0.58–0.64 in Hw; length of metathoracic femur 1.9–2.1; metathoracic tibia 2.1–2.2; length of S9+10 in lateral view 1.3–1.4; total length of cercus (dorsal branch) in lateral view 0.74–0.86; length of ventrobasal process in lateral view 0.12–0.22.
Unknown.
(6 males, 1 female). Holotype male, BRAZIL. Bahia State: Camacan municipality, Reserva Particular do Patrimônio Natural (RPPN) Serra Bonita, collecting point “Alojamento” (BA 2012-001; 15°23’16”S, 39°33’58”W, 810 m a.s.l.), 27–29.II.2012, R.A. Carvalho leg. (DZRJ 0050); 1 male paratype, same data but first trickle on road beyond managers house (15°23’19”S, 39°33’57”W, 710 m a.s.l. [822 m]), 5.I.2013, C.M. Flint & O.S. Flint Jr. leg. (USNM); 2 male paratypes, same data but quarry, 20.II.2015 (USNM; MZSP ODO-492, DNA voucher ENT2857); 1 male paratype, same data but 26.II.2015 (USNM); 1 female paratype, same data but spring near telecommunication towers (15°23’02”S, 39°34’19”W; 850 m a.s.l. [880 m]), 25.XII.2013, T.W. Donnelly leg. (MNRJ ODO-0001; ex-TWD collection). Additional specimen (excluded from type series). 1 male (teneral), same data as holotype but trickles at upper water tank (15°23’03”, 39°34’19”W, 890 m a.s.l.), 23.XII.2012, C.M. Flint & O.S. Flint, Jr. leg. (USNM).
Noun in apposition in reference to the type locality in the complex of RPPNs Serra Bonita, in honor of this protected area and its managers, who have taken positive steps towards the preservation of Atlantic Forest remnants in the southern portion of the biodiversity-rich state of Bahia.
Expected to be a submontane forest species, most likely occurring at seepages similar to the mesohabitat occupied by the large montane species from southeastern Brazil (e.g., F. itatiaiae and F. cf. lucia). The larvae possibly are madicolous and should inhabit shallow streams running over rocky substrates covered by ferns and hornworts/liverworts forming small waterfalls (Fig.
Habitus and habitat of species of Forcepsioneura: (35) F. regua sp. nov., holotype (Brazil. RJ: Cachoeiras de Macacu, DZRJ 2071); (36) F. serrabonita sp. nov., holotype (Brazil. BA: Camacan, DZRJ 0050), (37) F. regua sp. nov., live individual at type locality at Reserva Ecológica de Guapiaçu (REGUA); (38) overview of type locality at RPPN Serra Bonita; (39) shallow muddy and slow stream at Itatiaia National Park, typical habitat of small light blue species; (40) seepage covered by hornworts at Três Picos State Park, typical habitat of large orange-black species. Scale bars = 1 mm. Photos: (37) TMFK; (38) RA Carvalho; (39–40) APP.
The “Reserva Particular do Patrimônio Natural Serra Bonita” (RPPN, Private Reserve of Natural Heritage) is a complex of protected areas that started from a small fragment bought in the 1990s. Now it covers about 7,500 ha between 300 and 1,080 m above sea level of primary and secondary forests maintained by Instituto Uiraçu (
Like Serra Bonita, also in Southeastern Brazil, the “RPPN Reserva Ecológica de Guapiaçu” originated from private efforts aiming to conserve the AF. This protected area shares a similar history with Serra Bonita, beginning in the 1990s. Originally it was a farm and slowly it became a conservation and restoration portion of the AF. The preserve was founded in the beginning of 2000 and was officially recognized as a RPPN at the state level in 2013, at 302 ha. In 2014 a smaller, 35 ha area was incorporated to it and baptized as REGUA II. Unofficially, the preserve is between 6,500 and 7,500 ha, and 49% to 80% of its area is estimated to be inserted within the Três Picos State Park (
Recent studies highlight the relevance of REGUA. For example, 450 species of birds were recorded in REGUA and in neighboring areas (
These recent findings, including the two new species described here, bring to attention the urgency in directing collecting efforts at unexplored areas, and also the importance of private preserves that harbor the type localities of described species in the AF. Together, REGUA and Três Picos harbor over 210 species of Odonata, or about 24% of the known Brazilian dragonflies, and represent a major hotspot for the conservation of these insects. As highlighted by
When Forcepsioneura was erected,
Erected for Amazonian species (
The two species described here contribute at different levels to challenge the definition of Forcepsioneura. In F. serrabonita sp. nov. the posterolateral margin of the median lobe is projected into a well-defined tubercle-like process (Figs
The strong separation between AF and Amazonian species, somewhat resembling a vicarious pattern, is questionable. There are records of F. sancta (a typical AF species) from the Cerrado domain of the central Brazilian plateau (
Based on size, general coloration, shape of cercus and habitat, Forcepsioneura can be divided into two species groups: (1) includes F. garrisoni, F. haerteli, F. regua sp. nov. and F. sancta. Small, lowland species that present pale areas predominantly bluish-green, cercus slender with ventrobasal process comparatively long, and distribution almost restricted to coastal areas of the Atlantic Forest (F. sancta is an exception), some species even occupying strictly restinga formations, being associated with small shallow streams and muddy pools with slow running water (Fig.
The light blue Forcepsioneura form a complex of strongly similar species including F. garrisoni, F. regua sp. nov. and an undescribed species from southern state of Bahia. Although nothing was mentioned about blue hues in the original description of F. garrisoni (see
Additionally,
Finally, this paper is a first step towards an understanding of the diversity and morphology of this remarkable group of damselflies typical of the threatened Atlantic Forest domain. Studies on the morphological and spatial evolution of Forcepsioneura are being conducted.
This paper was only possible due to the help of Thomas W. Donnelly, Oliver Flint Jr (USNAM), Rosser W. Garrison (CDFA), and Angelo B.M. Machado (UFMG), who lent us specimens. We also thank ICMBIO/SISBIO for collecting permits; APP is strongly indebted to Rachel A. Carvalho, a colleague and specialist in leafhoppers who collected one of the holotypes and has shared interesting material from her expeditions; the protected area managers and employees, Nicholas and Raquel Locke and Jorge Bizarro (REGUA) and Vitor Becker (Serra Bonita) for their support of scientific research as a way to promote biological conservation. This study was partially supported by the following postdoctoral grants to APP: Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq-PDJ proc. 157592/2015-4) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES-PVE proc. A019/2013).